Stratégies dévelopmentales chez les larves de Calliphoridae : entre régulation thermique et socialité

On a cadaver, necrophagous dipteran larvae suffer from strong selection pressures during their development. The premise of this thesis is that such an extreme, competitive and constraining environment would have favored the emergence of efficient developmental strategies, based on mechanisms such as thermal regulation but also sociality. This PhD work is divided into three parts. The first part focuses on the thermal behavior of Calliphorids larvae, which are confronted with a heterogeneous thermic environment on the corps, in which they select the most appropriate area for their metabolic activity. Firstly, this part shows that larvae have a preferential developmental temperature, which is different according to the species (Lucilia sericata, Calliphora vomitoria and Calliphora vicina), although they exploit the same resource at the same time. Secondly, this part demonstrates that the larvae are always in search of this preferential temperature and thirdly, that they adapt both their displacement and their food intake according to the temperature of the nourishing substrate. This first part of experiments demonstrates that the temperature parameters have a strong effect on larval behavior. The second part of this work focuses on the social dimension of larval behavior by analyzing the influence of congeners, mainly through their active aggregation behavior. We show for Lucilia sericata a strong attractive and retentive effect of the group, making obvious that sociality prevails over thermal regulation. However, these results are radically different under heterospecific conditions where the group formation strongly depends on preferential temperatures as well as aggregation kinetics of each species. Finally, the third part of this work analyzes the effects of temperature and congeners on the development of individuals and shows that both the behavior of thermoregulation and the action of congeners impact the temperature selected by larvae, and therefore, their development. These results indicate the existence of individual and collective behavioral development strategies based on the optimization of multiple parameters that allow larvae to develop ideally in this extreme ecosystem of a decaying corpse.Les larves de Diptères nécrophages se développant sur un cadavre font face à de fortes pressions de sélection. Nous démontrons comment cet environnement extrême aurait favorisé l’apparition de stratégies comportementales efficaces et originales, basées sur des mécanismes comme la régulation thermique mais également la socialité.Ce travail pose en premier lieu les bases du comportement de régulation thermique des larves de Diptères Calliphoridae. En effet, celles-ci sont confrontées à un environnement thermique très hétérogène, dans lequel elles vont sélectionner la zone la plus appropriée à leur activité métabolique. Bien que différentes espèces exploitent la même ressource au même moment, nous avons observé que les larves de Lucilia sericata, Calliphora vomitoria et Calliphora vicina ont chacune une température préférentielle de développement. De plus, nous démontrons que ces larves sont en recherche constante de leur température préférentielle (thermorégulation), et qu’elles adaptent leur alimentation à la température du milieu. Ce premier volet d’expérimentations illustre ainsi le rôle prépondérant de la température dans le comportement des larves. En second lieu, ce travail s'intéresse à la dimension sociale des larves nécrophages, et plus particulièrement au comportement d'agrégation. Nous avons démontré chez Lucilia sericata un fort effet attractif et rétentif des congénères, rendant manifeste une prévalence de la socialité sur la régulation thermique. Les résultats sont cependant drastiquement différents dans des conditions hétérospécifiques, où la formation du groupe varie selon les températures préférentielles et les cinétiques d’agrégation de chaque espèce. Ainsi, la température sélectionnée par un groupe hétérospécifique émerge d'un compromis entre les comportements de thermorégulation et d'agrégation. Enfin, ce travail analyse l’effet de ces stratégies comportementales sur le développement des individus.Nous montrons que le comportement de thermorégulation et l’action des congénères affectent la température sélectionnée par les larves, et donc, leur développement. De tels résultats démontrent l’existence de véritables stratégies comportementales individuelles et collectives de développement, reposant sur l’optimisation de paramètres multiples permettant aux larves de se développer au mieux dans cet écosystème extrême qu’est le cadavre en décomposition

Developmental strategies of Calliphoridae larvae : between thermal regulation and sociality

Les larves de Diptères nécrophages se développant sur un cadavre font face à de fortes pressions de sélection. Nous démontrons comment cet environnement extrême aurait favorisé l’apparition de stratégies comportementales efficaces et originales, basées sur des mécanismes comme la régulation thermique mais également la socialité.Ce travail pose en premier lieu les bases du comportement de régulation thermique des larves de Diptères Calliphoridae. En effet, celles-ci sont confrontées à un environnement thermique très hétérogène, dans lequel elles vont sélectionner la zone la plus appropriée à leur activité métabolique. Bien que différentes espèces exploitent la même ressource au même moment, nous avons observé que les larves de Lucilia sericata, Calliphora vomitoria et Calliphora vicina ont chacune une température préférentielle de développement. De plus, nous démontrons que ces larves sont en recherche constante de leur température préférentielle (thermorégulation), et qu’elles adaptent leur alimentation à la température du milieu. Ce premier volet d’expérimentations illustre ainsi le rôle prépondérant de la température dans le comportement des larves. En second lieu, ce travail s'intéresse à la dimension sociale des larves nécrophages, et plus particulièrement au comportement d'agrégation. Nous avons démontré chez Lucilia sericata un fort effet attractif et rétentif des congénères, rendant manifeste une prévalence de la socialité sur la régulation thermique. Les résultats sont cependant drastiquement différents dans des conditions hétérospécifiques, où la formation du groupe varie selon les températures préférentielles et les cinétiques d’agrégation de chaque espèce. Ainsi, la température sélectionnée par un groupe hétérospécifique émerge d'un compromis entre les comportements de thermorégulation et d'agrégation. Enfin, ce travail analyse l’effet de ces stratégies comportementales sur le développement des individus.Nous montrons que le comportement de thermorégulation et l’action des congénères affectent la température sélectionnée par les larves, et donc, leur développement. De tels résultats démontrent l’existence de véritables stratégies comportementales individuelles et collectives de développement, reposant sur l’optimisation de paramètres multiples permettant aux larves de se développer au mieux dans cet écosystème extrême qu’est le cadavre en décomposition.On a cadaver, necrophagous dipteran larvae suffer from strong selection pressures during their development. The premise of this thesis is that such an extreme, competitive and constraining environment would have favored the emergence of efficient developmental strategies, based on mechanisms such as thermal regulation but also sociality. This PhD work is divided into three parts. The first part focuses on the thermal behavior of Calliphorids larvae, which are confronted with a heterogeneous thermic environment on the corps, in which they select the most appropriate area for their metabolic activity. Firstly, this part shows that larvae have a preferential developmental temperature, which is different according to the species (Lucilia sericata, Calliphora vomitoria and Calliphora vicina), although they exploit the same resource at the same time. Secondly, this part demonstrates that the larvae are always in search of this preferential temperature and thirdly, that they adapt both their displacement and their food intake according to the temperature of the nourishing substrate. This first part of experiments demonstrates that the temperature parameters have a strong effect on larval behavior. The second part of this work focuses on the social dimension of larval behavior by analyzing the influence of congeners, mainly through their active aggregation behavior. We show for Lucilia sericata a strong attractive and retentive effect of the group, making obvious that sociality prevails over thermal regulation. However, these results are radically different under heterospecific conditions where the group formation strongly depends on preferential temperatures as well as aggregation kinetics of each species. Finally, the third part of this work analyzes the effects of temperature and congeners on the development of individuals and shows that both the behavior of thermoregulation and the action of congeners impact the temperature selected by larvae, and therefore, their development. These results indicate the existence of individual and collective behavioral development strategies based on the optimization of multiple parameters that allow larvae to develop ideally in this extreme ecosystem of a decaying corpse

Les larves de diptères nécrophages en entomologie médico-légale : une histoire de température

L’entomologie médico-légale est l’utilisation des insectes afin de calculer l’intervalle post-mortem (temps écoulé entre la découverte d’un corps et la mort réelle de l’individu). Connaître la biologie, la physiologie et le comportement de ces insectes sur l’écosystème représenté par le cadavre est d’une importance primordiale pour améliorer les expertises entomologiques. C’est sur quoi se focalise l’unité de taphonomie médico-légale, unique équipe de recherche en France experte sur ce sujet

The maggot, the ethologist and the forensic entomologist: Sociality and thermoregulation in necrophagous larvae

Necrophagous insects are mostly known through forensic entomology. Indeed, experimental data investigating the effect of temperature on larval development underlies post-mortem interval estimations. However, such developmental studies rarely considered the behavior of maggots. In contrast, previous results supposed that calliphoridae larvae use behavioral strategies to optimize their development on carcasses. To test this idea, we analyzed the trade-off between thermal regulation (individual thermal preferences) and social behavior (aggregation) in Lucilia sericata larvae. The first set of experiments analyzed the behavior of third instars in response to thermal changes in their environment. The results demonstrated a clear thermoregulation behavior, supporting the assumption that larvae continuously move to reach a suitable internal temperature. The second set of experiments focused on the trade-off between thermal optimization and aggregation. The results showed a constant search for congeners and an attractiveness of aggregates, sometimes to the detriment of thermal optimization. Together, these results demonstrate a balance between behavioral thermoregulation and social strategies, two significant mechanisms for developmental optimization in necrophagous larvae. In conclusion, these findings highlights unexpected (social) strategies to cope with ephemeral resource and high selection pressure. They also raise important questions for forensic entomology. Keywords: Allee effect, Fitness, Maggot mass, Harsh environment, Trade-off, Blowflie

Thermoregulation in gregarious dipteran larvae: evidence of species-specific temperature selection

Due to the ephemeral nature of carcasses they grow on, necrophagous blowfly larvae should minimize the time spent on the cadaver. This could be achieved by moving to high-temperature areas. On that basis, we theorized that larvae placed in a heterogeneous thermal environment would move to the higher temperature that speed up their development. This study was designed to (1) test the ability of necrophagous larvae to orientate in a heterogeneous thermal environment, and (2) compare the temperatures selected by the larvae of three common blowfly species: Lucilia sericata (Meigen), Calliphora vomitoria (L.) and Calliphora vicina (Robineau-Desvoidy). For this purpose, we designed a setup we named Thermograde. It consists of a food-supplied linear thermal gradient that allows larvae to move, feed, and grow in close-to-real conditions, and to choose to stay at a given temperature. For each species and replication, 80 young third instars were placed on the thermal gradient. The location of larvae was observed after 19 h, with fifteen replications per species. The larvae of each species formed aggregations that were always located at the same temperatures, which were highly species-specific: 33.3 ± 1.52 °C for L. sericata, 29.6 ± 1.63 °C for C. vomitoria, and 22.4 ± 1.55 °C for C. vicina. According to the literature, these value allows a fast development of the larvae, but not to reach the maximum development rate. As control experiments clearly demonstrate that larval distribution was not due to differences in food quality, we hypothesized that the local temperature selection by larvae may result from a trade-off between development quality and duration. Indeed, temperature controls not only the development rate of the larvae, but also the quality of their growth and survival rate. Finally, results raise questions regarding the way larvae moved on the gradient and located their preferential temperature.SCOPUS: ar.jFLWINinfo:eu-repo/semantics/publishe

Mixed-species aggregations in arthropods

This review offers the first synthesis of the research on mixed-species groupings of arthropods and highlights the behavioural and evolutionary questions raised by such behaviour. Mixed-species groups are commonly found in mammals and birds. Such groups are also observed in a large range of arthropod taxa independent of their level of sociality. Several examples are presented to highlight the mechanisms underlying such groupings, particularly the evidence for phylogenetic proximity between members that promotes cross-species recognition. The advantages offered by such aggregates are described and discussed. These advantages can be attributed to the increase in group size and could be identical to those of non-mixed groupings, but competition-cooperation dynamics might also be involved, and such effects may differ between homo- and heterospecific groups. We discuss three extreme cases of interspecific recognition that are likely involved in mixed-species groups as vectors for cross-species aggregation: tolerance behaviour between two social species, one-way mechanism in which one species is attractive to others and two-way mechanism of mutual attraction. As shown in this review, the study of mixed-species groups offers biologists an interesting way to explore the frontiers of cooperation-competition, including the process of sympatric speciation